fossil soldier beetles from baltic amber of the anders ... filefabrizio fanti, anders leth damgaard...

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Baltic J. Coleopterol. 18(1) 2018ISSN 1407 - 8619

INTRODUCTION

The family Cantharidae, with more than 5,000species (Bouchard et al. 2009) and around 150genera, is widespread all over the world and inall continents except Antarctic, including smalloceanic islands, mountain areas, deserts(Konstantinov et al. 2009) and cold areas such

Fossil soldier beetles from Baltic amber of the Anders Damgaardamber collection (Coleoptera Cantharidae)

Fabrizio Fanti, Anders Leth Damgaard

Fanti F, Damgaard A. L. 2018. Fossil soldier beetles from Baltic amber of the AndersDamgaard amber collection (Coleoptera Cantharidae). Baltic J. Coleopterol. 18(1): 1 - 32.

In this paper, the authors describe and illustrate four new fossil genera and fifteen newfossil species of soldier beetles from Baltic amber belonging to the collection of AndersDamgaard: Eridanula n. gen. , Noergaardia n. gen. , Juratelacrima n. gen. ,Palmnickeneoceras n. gen., Cacomorphocerus bentifabrici n. sp., Cacomorphocerusmadseni n. sp., Eridanula susannaebierae n. sp., Noergaardia dinae n. sp., Cantharis(Cyrtomoptila) mikkelsenorum n. sp., Juratelacrima ballingi n. sp., Palmnickeneocerasejersboi n. sp., Podistra (Absidia) kloevedali n. sp., Rhagonycha (s. str.) nielsenae n.sp., Themus (Haplothemus) bennyianderseni n. sp., Malthinus (s. str.) rifbjergi n. sp.,Malthodes (s. str.) henningseni n. sp., Malthodes (s. str.) moellehavei n. sp., Kuskaellabajerae n. sp., and Autosilis annisettaekoppelae n. sp. The presence of the genus ThemusMotschulsky, 1858 in Eocene Baltic amber allows assumptions to be made about the originof the genus or the age of amber. The genera Eridanula and Norgaardia belong to therecently established tribe Cacomorphocerini Fanti & Kupryjanowicz, 2018 based on havingthe same shape of the antennae but with more antennomeres (17-19). Furthermore, Podistrakloevedali is the first described fossil species of the genus, until now known only at thegeneric level.

Key words: Cantharidae, new genera, new species, paleoentomology, Baltic, amber.

Fabrizio Fanti, via del Tamburino 69, I - 53040 Piazze (Siena), Italy. E-mail:[email protected]

Anders Leth Damgaard, Hornshøjparken 79, DK-7500 Holstebro, Denmark. E-mail:[email protected]

as North Greenland (Böcher 1995) andNorthern Alaska (Matthews & Telka 1997). Itis quite common even as fossils, 53 speciesbeing recently recorded in a catalogue (Fanti2017a) and 12 species subsequently described(Fanti 2017b, 2017c; Fanti & Castiglione 2017;Fanti & Kupryjanowicz 2017, 2018; Fanti &Vitali 2017; Fanti et al. 2018; Fanti & Michalski

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Fanti F, Damgaard A. L.

2018; Fanti & Pankowski 2018). The vast ambercollection of fossil soldier beetles of AndersDamgaard (Holstebro, Denmark) has alreadyprovided two new genera and species fromBurmese amber (Fanti et al. 2018). Therefore,the purpose of this work is to study thenumerous taxa from Baltic amber. Thecollection has proven to be particularly rich inspecies and also has two new genera related tothe newly established tr ibe of theCacomorphocerini Fanti & Kupryjanowicz,2018. Therefore, the Eocene subtropicalforests, as well as the current ones of SoutheastAsia and South America, were populated by manyspecies and genera of Cantharidae, with manyinteresting Eocene lineages now believed to beextinct.

MATERIALS AND METHODS

The amber specimens come from a quarry nearYantarny in the Kaliningrad region, Russia.Baltic amber is traditionally considered to beof Eocene age: Middle Eocene (Lutetian) (47.8-41.2 MY) to Late Eocene (Priabonian) (37.8-33.9 MY) but sometimes also Oligocene (e.g.Noetling 1883; Jeannel 1949; Laurentiaux1953; Vitali & Damgaard 2016). All amberspecimens were re-polished in order to highlightdorsal and ventral views, examined with a NikonSMZ 745T stereomicroscope and photographedwith a camera Imaging Source DFK 72AUC02attached to a trinocular microscope. Theholotypes are housed in the amber collectionof Anders Damgaard (ALDC) and subsequentlywill be deposited in the Zoological Museum,University of Copenhagen, Denmark (ZMUC).Subfamilies, tribes, genera and species arelisted in systematic alphabetical order.

SYSTEMATIC PALEONTOLOGY

Class Insecta Linnaeus, 1758Order Coleoptera Linnaeus, 1758Superfamily Elateroidea Leach, 1815Family Cantharidae Imhoff, 1856 (1815)

Subfamily Cantharinae Imhoff, 1856 (1815)Tribe Cacomorphocerini Fanti &Kupryjanowicz, 2018Genus Cacomorphocerus Schaufuss, 1892

Cacomorphocerus bentifabrici n. sp.(Fig. 1ABC)

Holotype. Female, in Baltic amber, accessionNo. ALDC0057/ALD.Ba.Can.8

Type locality. Russia, Kaliningrad Region,Sambian Peninsula, amber quarry near Yantarny(previously known as Palmnicken).

Type horizon. Middle Eocene (Lutetian) (47.8-41.2 MY) to Late Eocene (Priabonian) (37.8-33.9 MY).

Differential diagnosis. The genusCacomorphocerus is known for three speciesin Baltic amber: C. cerambyx Schaufuss, 1892,C. jantaricus (Kuśka & Kania, 2010) and C.wiszniewskii Fanti & Kupryjanowicz, 2018 andcited, at the generic level, also for the coevalRovno amber (Kazantsev & Perkovsky 2014).It is characterised by antennae 12-segmentedwith antennomeres III-IX dilated and saucer-shaped. This genus seems to be very uniform inthe habitus and only the pronotal shape seemsto be, in some cases, quite diagnostic. In fact,C. cerambyx shows a quadrate pronotum, withapical margin narrower, all angles roundish andsurface with longitudinally impression, while C.jantaricus shows a rectangular pronotum,concave to the sides, with basal angles slightlyprotruding laterally and a small denticle near theapical angles. Furthermore, the surface is roughwith deep punctation and two oblong tubercles(Schaufuss 1892; Kuśka & Kania 2010).Finally, C. wiszniewskii shows a subrectangularpronotum with concave sides from after the baseto the apical third and with a small denticle. Thenew species herein described, instead, has apronotum slightly longer than wide, with apicalmargin and strongly rounded sides in theanterior part, and the surface is rough with twooblong thickenings in the middle. In addition,C. bentifabrici n. sp. is entirely dark brown,

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Fossil soldier beetles from Baltic amber of the Anders Damgaard amber collection (Coleoptera Cantharidae)

Fig. 1. ABC: Cacomorphocerus bentifabrici n. sp. in Baltic amber. A: Holotype, dorso-lateralview, bar = 1.0 mm; B: Holotype, detail of head and pronotum, bar = 300 μm; C: Drawing ofpronotum, bar = 500 μm; D: Cacomorphocerus madseni n. sp. in Baltic amber. Drawing ofpronotum, bar = 500 μm.

while C. jantaricus has black-brown elytra withyellow elongate spots on humeri and pale brownmargin at apex (Kuśka & Kania 2010), C.cerambyx is probably entirely black (Schaufuss1892) and C. wiszniewskii is dark brown withblackish sternites (Fanti & Kupryjanowicz2018).

Description. Adult, winged, robust and stockyhabitus, female, entirely dark brown. Bodylength 8 mm (from front margin to elytral apex).Head rounded, very slightly narrower thanpronotum and posteriorly strongly covered bypronotum, surface with shallow punctation.Eyes mediocre, roundish, strongly prominent.Maxillary palps 4-segmented with lastpalpomere securiform. Antennae short, fewpubescent, reaching only the basal third of theelytra, 12-segmented, inserted in the inneranterior part at short distance from the eyes;scape elongated, stout, club-shaped; pedicel veryshort and cylindrical, about one-third as long asscape; antennomeres III-VI stout, slightly

saucer-shaped and asymmetrically inflated;antennomere VII saucer-shaped; antennomereVIII strongly saucer-shaped and asymmetricallydilated; antennomere IX slightly narrower thanantennomeres VII-VIII, subquadrate;antennomeres X-XII elongated and filiform.Pronotum slightly longer than wide; surface withsetae and shallow punctation; angles roundish;lateral and particularly the apical marginstrongly round and protruding to covering thehead, two evident and parallel thickenings in theposterior half and close to the basal margin,which is flat. Scutellum triangular, elongated andwith roundish apex. Elytra long, surpassing thelast abdominal segments, parallel-sided, withrounded apex, as wide as the pronotum, coveredwith numerous and short setae and with rugosesurface. Posterior wings covered by elytra.Sternum and abdomen pubescent (covered bymilky emulsion); last ventrite wide and nottriangular. Legs slender and fairly short; coxaemassive; trochanters elongated; femora slightlyenlarged and cylindrical with apex thin and

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completely fitted with short pubescence; tibiaewith pubescence, cylindrical slightly longer thanfemora. Tarsi 5-segmented, short and massive;first tarsomere elongated; second about 1.5times shorter than first tarsomere; third andfourth tarsomere triangular and shorter thansecond; fifth elongated and thin; claws simple,not well visible but probably with a delicatedenticle at base.Etymology. This new species is named inhonour of the Danish musician Bent Fabricius-Bjerre, internationally known as Bent Fabric, inrecognition of his career as a pianist andcomposer.

Syninclusions. Numerous wood remains (alsobotanical masses), stellate trichomes, and someair bubbles.

Remarks. The specimen is a female based onthe relatively short antennae compared withother Cacomorphocerus species, andparticularly for the last ventrite that is large andnot triangular.

The piece of amber (that measures 37 x 28 x 9mm) is not particularly transparent (lacquered).The inclusion is ventrally strongly covered by awhite cotton-like cloud (emulsion, milkysubstance) making it impossible to get a clearvision of the ventral side of the beetle.

Cacomorphocerus madseni n. sp.(Figs. 1D - 2)

Holotype. Male, in Baltic amber, accession No.ALDC0390/ALD.Ba.Can.15



Fig. 2. Cacomorphocerus madseni n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 1.0 mm;B: Holotype, ventral view, bar = 1.0 mm; C: Holotype, detail of pronotum, bar = 500 μm.

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Differential diagnosis. Cacomorphocerusmadseni n. sp. has the pronotum with deeptransversal incision, not present in any otherCacomorphocerus or fossil soldier beetles(Cacomorphocerus cerambyx Schaufuss, 1892has the longitudinal incision). It is alsocharacterised by the extremely shorts protibiaeand the pronotum with two strong expansionsat sides.

Description. Adult, winged, robust and stockyhabitus. Male, based on the triangular and narrowlast ventrite. Body length 6 mm. Entirelyblackish. Head slightly wider than pronotum,rounded, posteriorly covered by pronotum. Eyesroundish, strongly prominent. Maxillary palps4-segmented with last palpomere securiform.Labial palps 3-segmented with the last segmentelongated and slightly globular and pointed.Antennae short, slightly surpassing the humeri,12-segmented; scape elongated, stout; pedicelextremely short and roundish; antennomeres III-VI stout, slightly saucer-shaped andasymmetrically inflated; antennomeres VII-VIIIstrongly dilated and saucer-shaped, withantennomere VIII wider than antennomere VII;antennomere IX slightly narrower thanantennomere VIII; antennomeres X-XIIelongated and filiform. Pronotum convex, aswide as elytra, with in the middle a deep andsinuous transversal incision; basal marginstraight; apical margin slightly sinuous; sideswith two strong denticles - expansions roundedat apex, one near the basal angles and anotherone just above the median incision; surface withpunctation. Scutellum very wide, elongated, withroundish apex. Elytra very long, surpassing thelast abdominal segments, parallel-sided, withrounded apex, covered with numerous shortsetae. Posterior wings completely covered byelytra. Sternum and abdomen pubescent, withlast ventrite triangular and very narrow. Legsshort, slender, only the posterior long; coxaemassive; trochanters triangular, very narrow;femora slightly enlarged fitted with shortpubescence; protibiae extremely short, meso-and metatibiae long and cylindrical withpubescence. Tarsi 5-segmented; first tarsomere

elongated; second about 2.5 times shorter thanfirst tarsomere; third tarsomere triangular;fourth discoidal with anterior margin straight;fifth elongated and thin; claws simple, not wellvisible but seemingly with a denticle at base.

Etymology. This new species is named inhonour of the Danish novelist Svend ÅgeMadsen, in recognition of his writings.

Syninclusions. Stellate trichomes, a big airbubble and a few small wood remains.

Remarks. The piece of amber is transparent,roundish drop-shaped and measures 15 x 11 x 5mm. The legs of the inclusion are present, butsome of them are curled up and the rightmesotibia is broken and detached almost at thebeginning. The head and partially the pronotumare covered by a white emulsion, and thepronotum is also damaged. The last article ofthe right antenna is missing.

Genus Eridanula n. gen.

Type species. Eridanula susannaebierae n. sp.The genus is at present monotypic.

Etymology. The genus name derives fromEridanos (Greek = Eridanσs; Latin = Eridanus),derived from Ancient Greek “Hριδανός““Amber” + the Latin suffix -ula (diminutive) =small. This epithet refers to the name of themajor r iver of Baltica: Fenno-Sarmatia(Eridanos River), which carried and deposed theBaltic amber (Bijlsma 1981; Overeem et al.2001; Gibbard & Lewin 2016). Gender:feminine.

Diagnosis. The unequal maxillary palpomeres,with the last segment securiform and thepronotum without modified lateral margins,places this new genus in the subfamilyCantharinae. Eridanula n. gen. is characterisedby long elytra covering the abdomen; pronotumslightly longer than wide, with parallel lateralmargins and narrower than elytra; head globular,very large, dorsally telescopic and with evident

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Fig. 3. Eridanula susannaebierae n. gen. et n. sp. in Baltic amber. A: Holotype, dorso-lateralview, bar = 1.0 mm; B: Holotype, lateral view, bar = 1.0 mm.

Fig. 4. Eridanula susannaebierae n. gen. et n. sp. in Baltic amber. A: Holotype, detail of head andpronotum, bar = 200 μm; B: Holotype, detail of antennae, bar = 400 μm; C: Holotype, detail of lasttergites, bar = 500 μm.


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big eyes; claws simple without denticle (with asmall denticle in the other related genera) andantennae with 17 articles: the antennomeres III-VIII dilated and slightly saucer-shaped (notably,the antennomere VIII is very saucer-shaped), theantennomere IX subquadrate and the otherarticles filiform and slightly enlarged (which isunknown in any extant and fossil genera).

Eridanula susannaebierae n. sp.(Figs. 3 - 4)

Holotype. Female, in Baltic amber, accessionNo. ALDC0396/ALD.Ba.Can.19Type locality. Russia, Kaliningrad Region,Sambian Peninsula, amber quarry near Yantarny(previously known as Palmnicken).

Type horizon. Middle Eocene (Lutetian) (47.8-41.2 MY) to Late Eocene (Priabonian) (37.8-33.9 MY).Differential diagnosis. Given the similaritywith the Cacomorphocerus lineage, the numberof antennomeres and the shape of antennae areimportant diagnostic characters (see the nextgenus in this article). The only genus with thesame number of antennomeres isMichalskantharis Fanti, 2017 which has thearticles filiform and inflated but unmodified,and the antennae are also longer. Furthermore,Michalskantharis shows a pronotum totallyconvex, with apical and basal margins raised, andhead not laterally enlarged (Fanti 2017c).

Description. Adult, winged, elongate. Female,defined on the basis of the shape of last sternitenot narrowed, and relative short antennae. Bodylength 6 mm. Entirely blackish with legs,particularly the posteriors, slightly paler. Headalmost completely exposed, wider thanpronotum, large as elytra at humeri, laterallyvery enlarged and globular, dorsally telescopic,ventrally long and extruded, surface slightlyrugose covered with very sparse and short setae.Eyes in the lateral-dorsal part of head,extremely wide, evident, strongly convex,roundish. Mandibles not well visible, seems verywide at base and thin at apex. Unequal maxillary

palps, 4-segmented, with the last palpomeresecuriform. Antennae 17-segmented, rathershort and reaching almost half of the elytra,equipped with short pubescence and inserted inthe upper and inner part of the eyes and not closeto the eyes; scape very robust, elongated andcylindrical; pedicel short; antennomeres III-VIIdilated and slightly saucer-shaped; antennomereVIII saucer-shaped and wide; antennomere IXshort but large and subquadrate; antennomeresX-XVI filiform slightly enlarged and notparticularly long, antennomere XVII filiform.Pronotum much narrower than elytra, slightlylonger than wide; lateral margins parallel;surface with shallow punctation and recumbentpubescence, gibbous in the basal part and flat inthe apical part; apical and basal margins ratherraised. Scutellum wide at base and elongatedapically, where it is roundish. Elytra very long,narrow, covering the last abdominal segments,parallel-sided; apex almost straight and slightlyrounded, surface smooth without evidentpunctation and covered with short setae.Posterior wings dark brown, almost completelycovered by elytra. Sternum and abdominalsegments dark brown - blackish with shortpubescence, last tergite very short and roundishand last ventrite not narrow. Legs of mediumlength, covered with dense and numerous setae;coxae massive, elongated and globular;trochanters elongated and rounded at apex;femora enlarged and slightly curved, pro- andmesofemurs about 1.3 times longer than tibiae,metafemora slightly longer than metatibiae;tibiae thin, cylindrical, short, apically equippedwith spur. Tarsi 5-segmented; first tarsomereelongated and cylindrical, about 1.7-1.8 timeslonger than second; second tarsomere short;third tarsomere shorter than second and moreenlarged; fourth strongly bilobed; fifthelongated and flat, thinner than others; clawssimple without denticle at base.

Etymology. This new species is named inhonour of the Danish film director Susanne Bier,to thank her for her films and to recognize herfor the awards she has earned.


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Syninclusions. Air bubbles, botanical mess,very small wood remains, and a small mite.

Remarks. The amber piece (measures 15 x 14x 6 mm) is extremely transparent and allows anaccurate dorsal view. Only the left side iscovered by a white cotton-like cloud (emulsion,milky substance).

Genus Noergaardia n. gen.

Type species. Noergaardia dinae n. sp. Thegenus is at present monotypic.

Etymology. This genus is named in honour ofthe Danish author, journalist, debater andscriptwriter Lise Nørgaard + the Latin suffix -ia, as congratulation for her 100th birthday(1917-2017) and in honour of her decades ofcontributions to the Danish cultural treasurechest. She is a pioneer in consumer journalism,has fought for women’s rights and equality and,with her strong personality, has influenced andimpacted Danish culture. Gender: feminine.

Diagnosis. Long elytra covering the abdomen,pronotum without modified lateral margins, legswith 5-segmented tarsi and with the pretarsalclaws equipped by small denticle, and unequalmaxillary palpomeres with the last segmentsecuriform, are typical of the subfamilyCantharinae. This genus is also characterised by19-segmented antennae, antennal articles III-VIII dilated and saucer-shaped, IX rectangular-elongated and the others filiform (unknown inany extant and fossil genera).

Noergaardia dinae n. sp.(Figs. 5 - 6)

Holotype. Sex undetermined, in Baltic amber,accession No. ALDC0055/ALD.Ba.Can.6



Differential diagnosis. Easily distinguishablefrom all other soldier beetles by the 19-segmented antennae with saucer-shapedsegments III-VIII. Other similar genera withdissimilar antennae are Palmnickeneoceras n.gen. (11 articles, antennomeres III-V dilated,antennomere VI subrectangular),Cacomorphocerus Schaufuss (12 articles, III-IX saucer-shaped), Sucinocantharis Kuśka &Kania (16 articles, III-X dilated to saucer-shaped) and Eridanula n. gen. (17 articles, III-VII dilated, VIII saucer-shaped, IX slightlydilated). Electronycha Kazantsev (15 filiformantennal articles and only the VI-IX swollen)and Michalskantharis Fanti (17 antennalarticles but filiform inflated and not modified)could also be added to this group (Schaufuss1892; Kuśka & Kania 2010; Kazantsev 2013;Fanti 2017c).

Description. Adult, winged, flattened, elongate.Sex undetermined. Body length 8.0-8.5 mm anddifficult to assess because the head is folded.Entirely dark brown-blackish. Head globular, notwell visible, almost completely exposed andequipped with some setae. Eyes rounded.Maxillary palpomeres unequal; first segmentvery short; second elongated and third alongalmost one-half as long as the second; the lastsecuriform and very rounded at apex, typical ofthe subfamily Cantharinae. Antennae 19-segmented; scape elongated and much enlarged;second very short, strongly globular and thesmallest of all; antennomeres III-VII dilated andsaucer-shaped; antennomere VIII dilated,saucer-shaped/trapezoidal and more elongatedof the articles III-VII; antennomere IXrectangular elongated and slightly rounded;antennomeres X-XIX filiform with the first fivemore swollen at apex; antennomere XIX aboutone-half as long as the previous. Pronotumstrongly transverse and slightly convex withparallel sides and apical margin very reflanged,shiny, hair less, and with very small notimpressed punctation. Scutellum triangular with


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Fig. 5. Noergaardia dinae n. gen. et n. sp. in Baltic amber. ABC: Holotype, dorsal views (differentangulations), bars = 1.0 mm; D: Holotype, lateral view, bar = 1.0 mm.

Fig. 6. Noergaardia dinae n. gen. et n. sp. in Baltic amber. Holotype, detail of antenna, bar = 1.0mm.


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Fig. 7. Cantharis (Cyrtomoptila) mikkelsenorum n. sp. in Baltic amber. A: Holotype, dorsal view,bar = 1.0 mm; B: Holotype, ventral view, bar = 1.0 mm; C: Holotype, lateral view, bar = 1.0 mm; D:Holotype, detail of elytra, bar = 1.0 mm.

Fig. 8. Cantharis (Cyrtomoptila) mikkelsenorum n. sp. in Baltic amber. Holotype, detail of claws,bar = 400 μm.


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the base enlarged and concave and the apexrounded. Elytra long, slightly exceeding the lastabdominal segment, as wide as the pronotum,rounded at apex but not narrowed, with thesurface very granulose and equipped by longsetae. Sternites dark brown and tergites sub-squared with the last segment small in the lobe-shape. Coxae massive; trochanters triangular-elongated shaped with apex rounded; femorastrongly curved and cylindrical-globular with themetafemora more tapered from mid-length andalmost twice as long as the mesofemora; tibiaevery fine and cylindrical with metatibiae verylong, twice as long as the mesotibiae. Tarsi 5-segmented equipped by setae; first segmentlong; second almost one-half as long as the first;third one-half as long as the first and not bilobedat sides; fourth elongated and rounded at apex;fifth thin, very elongated; claws simple withpretarsal claws equipped by a rounded denticle.

Etymology. The species is named in honour ofDina Pallari, mother of the first author (Fanti),for her constant support and encouragement ofhis research and entomological studies.

Syninclusions. A few air bubbles and woodremains, some pyrite.

Remarks. The amber piece measures about 31x 18 x 9 mm and is very clear, enabling anoptimal dorsal view of the inclusion.

Tribe Cantharini Imhoff, 1856 (1815)Genus Cantharis Linnaeus, 1758Subgenus Cyrtomoptila Motschulsky, 1860Cantharis (Cyrtomoptila) mikkelsenorum n.sp.(Figs. 7 - 8)




Differential diagnosis. Cantharismikkelsenorum n. sp. is characterised by the lastpalpomere elongated and little securiform,pronotum transverse subrectangular withsinuous lateral margins and with ripples in themiddle. The other Cantharis in Baltic amber -Cantharis (s. str.) sucinonigra Kuśka, 1992and C. (Cyrtomoptila) sucinokotejai (Kuśka,1996) - are much smaller (6 mm) and showdifferent coloration and pronotal shape (Kuśka1992, 1996; Kazantsev 2013). Other remainsof fossil Cantharis, known at an undeterminedlevel, come from Baltic amber, Randecker Maar(Germany) and Puy-Saint-Jean near Clermont-Ferrand (France), with other extant species fromthe Pleistocene of the United Kingdom.Furthermore, other species have been describedfrom the Early Miocene of Radoboj in Croatia,the Upper-Middle Miocene of Oeningen inGermany, and the Oligocene of Rott(Siebengebirge) in Germany (Fanti 2017a).

Description. Adult, winged, flattened. Male,defined on the basis of the shape of last sternite,which is short and vaguely triangular, and forthe aedeagus in part visible. Body length 10.5mm (prothorax to abdomen 8 mm, head ~2.5mm). Entirely blackish-dark brown with reddish-brownish legs. Head almost completelyexposed, covered with numerous small granulesand with long setae on the forehead. Eyes verylarge, rounded and strongly convex. Mandibleslong, black, without tooth and very sharp at apex.Maxillary palps with the last segment elongated,cylindrical, slightly enlarged in the lower partand rounded at apex. Antennae inserted very nearand in the upper part of the eyes, 11-segmented,short and not surpassing the elytral half,filiform, with thick interspersed setae and otherlonger; scape long, cylindrical, narrowed frommiddle to the base; pedicel short, one-third aslong as scape; antennomeres III-IV cylindricalfeebly inflated, as long as scape, antennomeresV-XI filiform subequal in length and very long.Pronotum transverse, subrectangular, equipped


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with a short pubescence and with blows, ripplesin the middle, with the reflanged basal marginand sinuous lateral margins. Scutellum reddish-brown, triangular with roundish base and pointedapex. Elytra wider than pronotum, long,surpassing the last abdominal segments, coveredby numerous setae, parallel-sided and withrounded apex. Posterior wings dark and foldedbetween the tergites and elytra. Sternum andabdominal segments dark-brown; last tergitelarge and rounded; last sternite small, vaguelytriangular. Legs covered with pubescence; coxaeshort and massive; trochanters triangular;femora enlarged, particularly the metafemorawhich are also curved; pro- and mesotibiae thinand cylindrical, ~1.5 times as long as femora;metatibiae elongated, ~1.7 times as long asmetafemora. Tarsi 5-segmented equipped withsetae; first segment slightly elongated andenlarged; second 0.7 times as long as the first;third evidently bilobed at sides; fourth segmenttriangular inverted with the tip toward the thirdsegment; fifth thin and elongated; claws simplewithout tooth at the base. Aedeagus extruded inpart.

Etymology. This new species is named inhonour of the Danish actors Lars DittmannMikkelsen and Mads Dittmann Mikkelsen, inrecognition of their contr ibution to thetelevision, theatre and film industries.

Syninclusions. Wood remains (of which twoare wooden frustules), stellate trichomes,pyrite, mite.

Remarks. The amber piece, with almost semi-circular shape, measures approximately 24 x 17x 5.5 mm. The new species herein described isinserted in the subgenus Cyrtomoptila for theclaws simple (Fig. 8), without tooth at the base(in males, however, the tooth tend to be verysmall but present, though sometimes missing).The rounded remains visible at the apex of thefifth tarsomere and on the dorsal part of the claware probably wood fragments.

Genus Juratelacrima n. gen.

Type species. Juratelacrima ballingi n. sp. Thegenus is at present monotypic.

Etymology. From the name of Goddess(sometimes described as a mermaid or undine)Jūratė (Jurata in Polish), of the Lithuanian (andBaltic) legend/tale, plus the Latin noun (inapposition)“lãcrima” = tear, crying, resin(understood here as a tear of Goddess Jūratė).The Goddess lived under the Baltic Sea in abeautiful amber castle, when a young fishermannamed Kastytis was disturbing the peace andcatching a lot of fish, and Jūratė decided topunish him but instead fell in love. Thus,Perkūnas, the thunder god, furious that animmortal goddess had fallen in love with amortal man, destroyed the amber castle (in othervariations Perkūnas kills Kastytis). Therefore,the amber pieces found would be the remainsof the castle or the tears of Jūratė, saddened bythe death of Kastytis. Gender: feminine.

Diagnosis. The new genus is characterised bythe third tarsomere straight at apex and notbilobed, claws simple with a small denticle atthe base, the pronotum gibbous, which isanteriorly not narrow and with straight sides andexpanded apical margin, and head slightlyelongated behind eyes. The unequal maxillarypalpomeres, with the last segment securiform,clearly make this new genus attributable to thesubfamily Cantharinae.

Juratelacrima ballingi n. sp.(Fig. 9)

Holotype. Male, in Baltic amber, accession No.ALDC0458/ ALD.Ba.Can.22




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Differential diagnosis. The new genus issimilar to Rhagonycha, from which it differsin the pronotum not anteriorly narrowed and inthe claws simple instead of bifid at the apex.The third tarsomere, straight at apex, notbilobed, is also present in the members of thetribe Podabrini Gistel, 1856, but they have thehead distinctly constricted behind the eyes andthe pronotum mostly truncate anteriorly(Pelletier & Hébert 2014). Furthermore, thefossil genus Sucinorhagonycha Kuśka, 1996has 12 antennomeres and claws simple, withoutdenticles or lobes (Kuśka 1996; Kubisz 2000;Fanti 2017a).

Description. Adult, winged, slender. Male,defined on the basis of the long antennae andlast sternite narrow. Body length ~7.2 mm; elytra6.0 mm; antennal length ~6.3-6.4 mm. Entirelyblackish. Head not completely exposed, widerthan pronotum, roundish, slightly elongatedbackwards the eyes, surface slightly rugose andwith small punctation and setae, frons concave.Eyes wide, lateral on head, rounded, coveringaround half of the temples, interocular distance~1.2 times greater than eye diameter. Mandiblessmooth without denticles, wide at base and thinat apex. Maxillary palps unequal, 4-segmented,with the last palpomere securiform. Labial palps3-segmented, with long articles and the lastpalpomere elongated and pointed. Antennae 11-segmented, filiform, very elongated, reachingalmost the apex of the elytra and almost the lastabdominal segment, inserted in proximity to theeye and in its central-anterior part; scape veryrobust, elongated and cylindrical, slightlyenlarged in the middle; pedicel thin, short and~2.3 times shorter than scape; antennomere IIIfiliform, elongated and 2.8 times longer thanpedicel; antennomeres IV-IX filiform, subequal,very elongated and slightly longer than third;antennomere X filiform, slightly shorter thanprevious; antennomere XI thin and as long asIV; all articles equipped with short setae on thesides. Pronotum narrower than elytra, longerthan wide; lateral margins parallel; surface witherected pubescence, gibbous with twolongitudinal thickenings in the basal part andparts near the apical and basal margins

transversely concave; apical margin extended.Scutellum triangular. Elytra elongated, coveringthe last abdominal segments, parallel-sided; apexrounded; surface smooth covered with erectedand scattered setae. Posterior wings dark brown,almost completely covered by elytra. Sternumand abdominal segments with short pubescence;last sternite and tergite elongated, narrow, withlong setae. Legs elongated, covered with denseand numerous setae; coxae massive, elongatedand globular; trochanters elongated; femoracylindrical, slightly enlarged, not curved; tibiaethin, cylindrical, slightly longer than femora.Tarsi 5-segmented; first tarsomere elongated,about 1.8 times longer than second; secondtarsomere straight at margin; third tarsomereshorter than second, not bilobed at sides; fourthshort, bilobed; fifth elongated, thinner thanothers; claws simple with small and obtuse lobeat base.

Etymology. This new species is named inmemory of the Danish film and TV director ErikBalling (Nyborg, 29 November 1924 - Gentofte,19 November 2005).

Syninclusions. Wood remains.

Remarks. The amber piece is very transparent,rounded and measures 24 x 22 x 6 mm. Theinclusion is complete, except for the leftantenna that is preserved until the antennomereII.

Genus Palmnickeneoceras n. gen.

Type species. Palmnickeneoceras ejersboi n.sp. The genus is at present monotypic.

Etymology. From Palmnicken (now Yantarny)the old name of the locality of the amberdeposits, the word-forming element “eo” =earliest, derived to Greek “eos” = dawn,morning, daybreak (in reference to Eocene, thesecond Epoch to the Tertiary Period) and“ceras”, the simple Latin transliteration of theGreek “keras” = horn, antenna. Gender: neuter.


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Fig. 9. Juratelacrima ballingi n. gen. et n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 1.0mm; B: Holotype, lateral view, bar = 1.0 mm; C: Holotype, detail of head and pronotum, bar = 500μm; D: Holotype, detail of tarsomere and claws, bar = 100 μm; E: Holotype, detail of palps, bar =100 μm.

Fig. 10. Palmnickeneoceras ejersboi n. gen. et n. sp. in Baltic amber. A: Holotype, dorsal view, bar= 500 μm; B: Holotype, ventral view, bar = 500 μm; C: Holotype, detail of antennae, bar = 500 μm;D: Holotype, dorsal view, bar = 1.0 mm.


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Diagnosis. The new genus is characterised bylong elytra covering the last abdominalsegments; pronotum transverse with parallelsides and basal margin slightly bordered; 11antennal articles; antennomeres III-V dilated/inflated and very slightly saucer-shaped (similarto Cacomorphocerus where the “central”antennomeres are saucer-shaped); antennomereVI subrectangular to slightly basally elongatedand the others filiform. Furthermore, themaxillary palps are very elongated and seemslightly securiform, which makes theassignment to the subfamily Cantharinaeuncertain and is here tentatively included in thetribe Cantharini.

Palmnickeneoceras ejersboi n. sp.(Fig. 10)




Differential diagnosis. Palmnickeneocerasejersboi n. sp. is distinguishable by the palpselongated and slightly securiform, and by theshape of antennae, unknown in any other fossilspecies. The other fossil genera of the samelineage show more antennomeres.Furthermore, extant and fossil members of thesubfamily Malthininae with long elytra,Malthinus (Indomalthinus) Brancucci inWittmer & Brancucci, 1978, Inmalthodes Pic,1938, Macrocerus Motschulsky, 1845,Mimomalthinus Pic, 1931, MantimalthinusFanti & Castiglione, 2017 and Kuskaella Fanti& Kupryjanowicz, 2017 - show a differentsurface of elytra, different palps or a differentpronotal shape (Fanti & Castiglione 2017; Fanti& Kupryjanowicz 2017).

Description. Adult, winged, robust habitus. Sexundetermined (last sternites are not wellvisible). Body length 5 mm. Pronotum reddish- orange, elytra dark brown blackish, antennaeand legs testaceous - brown. Head pale brown,as large as pronotum. Eyes small and elongated-subspherical, interocular dorsal distance ~4.2times greater than eye diameter. Mandiblessmall, little visible but seemingly without tooth.Maxillary palpomeres unequal with last segmentvery elongated and slightly securiform, withoutappendix at apex. Antennae inserted near and onthe inner part of the eyes, long, reaching theapical third of the elytra, 11-segmented,covered with dense and very short setae; scapeelongated and cylindrical; pedicel very short andabout six times shorter than scape;antennomeres III-V dilated; antennomere VIrectangular; antennomeres VII-XI filiform,antennomeres XI extremely elongated, muchmore than scape. Pronotum strongly transverse,parallel-sided; basal margin very slightlybordered; surface smooth with some smallsetae. Scutellum wide at base and elongated atapex, that is roundish. Elytra slightly wider thanpronotum with rugose surface and numeroussetae, elongated, covering the last abdominalsegments; apex slightly roundish. Posteriorwings covered by elytra. Sternum and abdominalsegments dark brown and covered with numeroussetae. Legs equipped with numerous short setae;coxae small and short; trochanteres elongatedand roundish; profemora curved, very dilated andcylindrical, meso- and metafemora slender andlonger; tibiae cylindrical very thin and long.Tarsi 5-segmented; first tarsomere slightlyelongated; second tarsomere about one-half aslong as the first, third tarsomere very small andbilobed at sides; fourth triangular; fifth long andscale-shaped; claws simple, long and robust.

Etymology. The species is named in memoryof the Danish journalist and writer Jakob Ejersbo(Rødovre, 6 April 1968, Aalborg, 10 July 2008).

Syninclusions. Numerous stellate trichomesand wood remains (also botanical masses andpollen grains), pyrite, one Trichoptera, twoDiptera, one mite.


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Remarks. The piece of amber of rectangularshape, measures 24 x 10 x 5 mm. The leftantenna is folded along the body, theantennomeres VIII-X are not well visible andeven all legs are curled up. It is not possible toobserve if the claws have a tooth at the base.

Genus Podistra Motschulsky, 1839Subgenus Absidia Mulsant, 1862

Podistra (Absidia) kloevedali n. sp.(Fig. 11)




Differential diagnosis. No fossil Podistra hasbeen described until now (Fanti 2017a).Podistra kloevedali n. sp. is tentativelyassigned to the subgenus Absidia for the clawswith tooth at the base, because claws are simplein the subgenera Pseudoabsidia Wittmer, 1969and Hemipodistra Ganglbauer, 1922. The newspecies is distinguishable from Cantharis (s.str.) sucinonigra Kuśka, 1992 by the larger size(10 mm instead of 6 mm), coloration, differentantennomeres, in particular, the pedicel almostthree times shorter than III (twice shorter thanIII in C. sucinonigra), and by the pronotumstraight laterally instead of rounded. It is alsodistinguishable from Cantharis (Cyrtomoptila)sucinokotejai (Kuśka, 1996), which is the mostsimilar species, by the different coloration, thelarger size and eyes, and by the claws with toothat base (without lobe or tooth in C.sucinokotejai). Finally, it differs fromCantharis (Cyrtomoptila) mikkelsenorum n.sp. in the different colour, claws, pronotumwithout ripples in the middle and straight atsides, and in the palps typically securiform.

Description. Adult, winged, flattened. Female,defined on the basis of the shape of last uritesthat are very wide and on short antennae. Bodylength 10 mm. Entirely reddish-dark brownexcept for the black head. Head completelyexposed, pubescent, elongated and covered withnumerous granules. Eyes extremely large,roundish and strongly convex. Mandibles notvisible. Maxillary palps 4-segmented with thelast palpomere securiform. Labial palps 3-segmented with the last palpomere elongatedand securiform. Antennae inserted very near andin the upper part of the eyes, 11-segmented,short and slightly surpassing the humeral zoneof elytra, filiform, with scattered long setae;scape robust, long and cylindrical; pedicel short,one-third as long as scape; antennomeres III-VIII filiform, subequal, and slightly shorter thanscape; antennomeres IX-XI filiform, thin,subequal in length and slightly shorter than theprevious. Pronotum longer than wide,rectangular (typical of Podistra), as wide as thehead and equipped by a short pubescence; basalsurface convex and middle-anterior part flat;margins bordered and sides straight; propleuraroundish. Scutellum reddish-brown, triangular,elongated and with slightly pointed apex. Elytramuch wider than pronotum, elongate, coveringthe last abdominal segments, equipped by longand sparse setae and several short setae, parallel-sided and with rounded apex. Posterior wingscovered by elytra. Abdominal segments largeand rectangular with short pubescence. Legs verypubescent; coxae short and massive; trochanterselongated and enlarged; femora robust; pro- andmesotibiae thin, short and cylindrical, equippedby two spurs, metatibiae very long. Tarsi 5-segmented; first segment slightly elongated andenlarged; second slightly shorter than first (inthe metatarsi the second is one-half length ofthe first); third segment bilobed at sides; fourthsegment strongly bilobed at sides with lobesthinner and longer than lobes of the thirdsegment; fifth thin and elongated; claws simplewith very thin and slightly elongated tooth at thebase (posterior claws simple, without denticle).

Etymology. This new species is named inhonour of the Danish pioneer, longtime sailor,


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Fig. 11. Podistra (Absidia) kloevedali n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 1.0mm; B: Holotype, detail of head and pronotum, bar = 500 μm; C: Holotype, detail of antenna, bar= 800 μm; D: detail of tergites and apex of elytra, bar = 1.0 mm.

Fig. 12. Rhagonycha nielsenae n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 500 μm; B:Holotype, ventral view, bar = 500 μm; C: Holotype, detail of pronotum, bar = 100 μm; D: Holotype,detail of sternum, bar = 200 μm.


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author and lecturer Troels Kløvedal (born TroelsBeha Erichsen), in recognition of his trips withthe Nordkaperen ship.

Syninclusions. Air bubbles, wood remains, oneDiptera, and one small leg (probably of midge).

Remarks. The rectangular amber piecemeasures 20 x 46 x 5.5 mm. The inclusion isventrally and laterally covered by a white cotton-like cloud (emulsion, milky substance).

Genus Rhagonycha Eschscholtz, 1830Subgenus Rhagonycha Eschscholtz, 1830

Rhagonycha (s. str.) nielsenae n. sp.(Fig. 12)




Differential diagnosis. Only two species ofRhagonycha are actually known from amberdeposits: Rh. kryshtofovichi (Yablokov-Khnzorian, 1960), which differs from Rh.nielsenae n. sp. in larger size (7 mm) anddifferent length of the antennomeres, and Rh.sucinobaltica Poinar & Fanti, 2016, whichdiffers in the blackish-dark brown colour, thepronotum with much more concave sides andin the different last abdominal segment(Yablokov-Khnzorian 1960; Kazantsev 2013;Poinar & Fanti 2016). Other members of thegenus Rhagonycha are known from Eocene ofFlorissant (USA), Miocene of Oeningen(Germany) and Radoboj (Croatia), and fromPliocene of Lac Chambon, France (Fanti2017a). All show different colour and body size.

Description. Adult, winged. Female, defined onthe basis of the short antennae, massive habitusand on the large last ventrite. Body length ~3.3mm; elytra 2.5 mm. Entirely dark brown exceptfor the black head. Head completely exposed,elongated, pubescent. Eyes large and roundish,interocular ventral distance about two timesgreater than eye diameter. Mandibles notvisible. Maxillary palps 4-segmented with thelast palpomere elongated-securiform and stout.Labial palps 3-segmented with the last segmentsecuriform. Antennae filiform, 11-segmented,short and slightly surpassing the humeral zoneof elytra, equipped with short setae; scaperobust, long and cylindrical; pedicel short, one-third as long as scape; antennomere III similarin length to the pedicel; antennomeres IV-VIvery elongated and subequal in length;antennomeres VII-XI subequal in length, shorterand more robust than the previous. Pronotumtrapezoidal, slightly wider than long and largerthan head, equipped by a dense short pubescenceand small punctation, large at base and graduallysteadily narrower towards the apex; basal anglesstrongly rounded; apical margin bordered.Scutellum strongly rounded and pubescent.Elytra elongated and covering the last abdominalsegments, stout, very slightly broader than baseof pronotum, parallel-sided; apex narrow andslightly rounded; surface equipped with manysetae. Posterior wings completely covered byelytra. Sternum dark brown, trapezoidal,equipped with scattered setae; ventrites shortand wide with the last rounded and large. Legsshort and pubescent; coxae massive; trochanterselongated with rounded apex; femora robust;tibiae thin, cylindrical, as long as femora. Tarsi5-segmented; first segment slightly elongated;second and third shorter than first and straightat apex; fourth segment slightly bilobed at sides;fifth thin and elongated; claws bifid but not wellvisible.

Etymology. This new species is named inmemory of Nielsine Mathilde Nielsen(Svendborg, 10 June 1850 - Copenhagen, 8October 1916), the first female academic andphysician in Denmark.


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Syninclusions. Wood remains.

Remarks. The amber piece is yellow and quiteflattened and measures around 15 x 9 mm. Theinclusion is complete with the ventral side notoptimally visible, and dorsally covered with asuperficial cotton-like emulsion.

Genus Themus Motschulsky, 1858Subgenus Haplothemus Wittmer, 1973Themus (Haplothemus) bennyianderseni n.sp.(Fig. 13)




Differential diagnosis. Large size and longcheeks are quite typical of the genus Themusand the rounded sides of the pronotum arepresent only in the subgenus Haplothemus(Švihla 2008). Themus bennyianderseni n. sp.is distinguishable from Th. (Haplothemus)pristinus Kazantsev, 2013, the only othercongener in amber, by sides and apical marginof the pronotum strongly rounded, without bluntand noticeable angles (Kazantsev 2013). Asindicated in Fanti (2017a), other Themus fossilsare known for the Middle Miocene of Shandong,Shanwang, Linqu County in China (Zhang 1989;Zhang et al. 1994).

Description. Adult, winged, very robust andstocky habitus. Sex undetermined. Body length15 mm; width at humeri ~3.5 mm. Real colourdifficult to determine but head and pronotumare probably bluish-grey, elytra dark testaceous-grey, knees and tibiae black. Head exposed andstrongly elongated, narrower than pronotum,

without punctation but with small granules andsome long setae, parallel-sides behind eyes, andwith long cheeks. Eyes roundish, small and veryconvex; interocular distance about 3.8 timesgreater than eye diameter. Mandibles short,massive and powerful. Maxillary palpomeresunequal; first segment very short; secondelongated and strongly globular; third small; lastsecuriform. Antennae 11-segmented, filiform,hairless except for a tuft at the apex of eachsegment, placed away from the apical part ofthe eyes toward the clypeus, posteriorly reachingthe metafemurs; antennomeres subequal inlength except for the second and third, whichare slightly shorter. Pronotum with very smalland not impressed granulate punctation, withshort setae, very rounded apical margins andsides (unknown in any extant and fossilcongeners), basal margin perfectly straight.Scutellum blackish, triangular with largelyrounded apex. Elytra long, very slightly broaderthan pronotum, slightly widened at the basalthird, with roundish apex, granulose with scarceand small raised setae. Posterior wingscompletely covered by elytra. Legs two-tone;trochanters short and roundish except for themetatrochanter, which is elongated; femorashort and massive with setae; tibiae relativelyfine and ~2 times longer than femora, equippedby numerous small and short setae; metatibiaecurved. Five tarsomeres; the first very elongatedand swelled after half-length; second one-halfas long as the first; third and fourth about two-thirds as long as the second; fifth tarsomere andclaws not well visible.

Etymology. This new species is named inhonour of the Danish songwriter, poet, author,composer and pianist Benny Andersen, inrecognition of his long, successful career.

Syninclusions. Wood remains, stellatetrichomes, some air bubbles, pyrite, someDiptera, parts of a spider.

Remarks. The rectangular (rounded at apex)piece of amber (lacquered) measures around 38x 27 x 12 mm, with the surface showing a


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shallow fracture that does not reach theinclusion. Right metatarsus is not conserved.

Subfamily Malthininae Kiesenwetter, 1852Tribe Malthinini Kiesenwetter, 1852Genus Malthinus Latreille, 1806Subgenus Malthinus Latreille, 1806

Malthinus (s. str.) rifbjergi n. sp.(Fig. 14)




Differential diagnosis. Fossil records ofMalthinus are known from Baltic amber asvarious specimens at the generic level (Fanti2017a) and only one described species, M.danieli Kuśka & Kania, 2010. Malthinusrifbjergi n. sp. differs in the brown legs,pronotum and elytra and the yellow head(pronotum yellow with a longitudinal black spoton the basal half, tibiae yellow-brown andfemora dark in M. danieli) (Kuśka & Kania2010). Furthermore, the new species has adifferent length of antennomeres, the secondarticle being as long as third, and fourth and fiftharticles longer, while in M. danieli the secondis shorter and the antennomeres III-XI aresubequal in length. The body length of M. danieliis 3.2 mm, while M. rifbjergi n. sp. is 4.0 mm.

Description. Adult, winged, slender. Female,defined on the basis of the short antennae andlarge last ventrite. Body length 4.0 mm. Headyellow and pronotum, elytra, antennae and legsbrown. Head completely exposed, stronglytriangular behind eyes, wider than pronotum,very wrinkly and equipped with impressed

punctation. Eyes large, round, convex. Maxillarypalps 4-segmented, with palpomeres unequal inlength, with the terminal globular, elongated anddistally pointed. Labial palps 3-segmented withthe last segment globular and pointed. Antennae11-segmented, filiform, surpassing the half ofelytra; scape very elongated, club-shaped andenlarged at apex; pedicel ~1.5 times shorter thanscape; antennomeres III-VI as long as pedicel;antennomeres VII-X slightly shorter than III;antennomere XI as long as III, slightly enlargedfrom middle to apex which is slightly pointed;all antennomeres equipped with many shortsetae. Pronotum transverse, slightly narrowedanteriorly, equipped with scattered pubescence,apical and basal margins straight and stronglybordered; sides sinuous; angles slightly rounded;surface not flat. Scutellum short and rounded.Elytra slightly narrower than head, wider thanpronotum, short and not covering the lastabdominal segments, parallel-sides, slightlynarrowed from half length; apex stronglyrounded; surface covered with impressedpunctation ordered in lines and equipped withsetae; apex without yellow rounded marks.Posterior wings dark, long and very surpassingthe elytra and the abdominal segments. Sternumand abdominal segments pubescent; last tergitevery short and rounded at apex. Legs slender;coxae massive; trochanters elongated; femoravery enlarged; tibiae cylindrical, thin, longerthan femora. Tarsal formula 5-5-5; tarsomere Ilong and ~2 times longer than second;tarsomere III short, tarsomere IV bilobed;tarsomere V slender; claws simple.

Etymology. This new species is named inmemory of the Danish writer Klaus Rifbjerg(Copenhagen, 15 December 1931 -Copenhagen, 4 April 2015).

Syninclusions. Wood remains, oakinflorescence, stellate trichomes, some Diptera(Empididae and other undetermined family).

Remarks. The amber piece measures 19.5 x14.5 x 9 mm and weighs 2.4 grams. Theinclusion is complete and well visible.


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Fig. 13. Themus (Haplothemus) bennyianderseni n. sp. in Baltic amber. A: Holotype, detail ofhead and pronotum, bar = 1.0 mm; B: Holotype, dorsal view, bar = 1.0 mm; C: Holotype, globalaspect of the amber specimen with the inclusion (this photo is also found in FANTI 2017a), bar =5.0 mm.

Fig. 14. Malthinus rifbjergi n. sp. in Balticamber. A: Holotype, dorsal view, bar = 1.0mm; B: Holotype, detail of head andpronotum, bar = 500 μm.

Tribe Malthodini Böving & Craighead,1931Genus Malthodes Kiesenwetter, 1852Subgenus Malthodes Kiesenwetter, 1852

Malthodes (s. str.) henningseni n. sp.(Fig. 15)

Holotype. Male, in Baltic amber, accessionNo. ALDC0459/ALD.Ba.Can.23

Type locality. Russia, Kaliningrad Region,Sambian Peninsula, amber quarry nearYantarny (previously known as Palmnicken).Type horizon. Middle Eocene (Lutetian)(47.8-41.2 MY) to Late Eocene(Priabonian) (37.8-33.9 MY).

Differential diagnosis. The habitus, sizeand lobe-shaped last tergite could make itsimilar to the M. brevicollis group (Liberti


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Fig. 15. Malthodes henningseni n. sp. in Baltic amber. A: Holotype, dorso-lateral view, bar = 200μm; B: Holotype, ventro-lateral view with detail of urites, bar = 200 μm; C: Holotype, detail ofurites, bar = 100 μm; D: Reconstruction of last abdominal segments, bar = 100 μm.

Fig. 16. Malthodes moellehavei n. sp. in Baltic amber. A: Holotype, dorso-lateral view, bar = 500μm; B: Holotype, detail of last abdominal segments, bar = 200 μm; C: Reconstruction of lastabdominal segments, bar = 100 μm.


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2016), but the last sternite shows a peculiarform, unknown in any fossil and CentralEuropean (and Italian) Malthodes.

Description. Adult, winged, robust. Male, basedon the last urites modified. Body length 2.5 mm.Entirely brown without yellow spots on elytra.Head completely exposed, strongly rounded andconvex, as wide as pronotum, equipped withscattered and strong punctation and with shortsetae. Eyes very large, round, convex and veryprominent. Maxillary palps 4-segmented,unequal in length, with the first small, secondelongated and apically enlarged, third robust,terminal one globular and distally very pointed.Labial palps 3-segmented with the last segmentglobular and pointed. Antennae filiform (theantennomeres are not completely preserved, butwe assume to be 11 articles as for all speciesof the genus), scape extremely elongated, club-shaped and swollen from half length to apex;pedicel filiform and about 1.7 times shorter thanscape; antennomeres III-VI filiform and robust;antennomeres pubescent. Pronotum transverse,equipped with a shallow punctation and scatteredpubescence; apical margin straight and slightlybordered; sides sinuous and bordered; anglesrounded; surface concave in the middle.Scutellum triangular. Elytra slightly wider thanpronotum, long and reaching the middle of uriteVII, wide at base and narrowed after half thelength with roundish apex, surface rugose andpubescent. Posterior wings wide, long andevidently surpassing the elytra and theabdominal segments, dark and equipped withsetae along the edges. Legs slender; coxaerobust and very elongated; trochanters roundishand only slightly elongated; femora enlarged;tibiae cylindrical, very thin, longer than femora.Tarsal formula 5-5-5; tarsomere I long androbust; second flat and shorter than first; thirdtriangular and about 1.2 times shorter thansecond; tarsomere IV bilobed; tarsomere Vslender; claws simple. Penultimate tergite (tg9)elongated and not particularly transverse; lasttergite (tg10) lobe-shaped and elongated withstraight apex; last sternite (st9) long, curvedfrom half length, apically flat and rounded and

with an appendix (lamina) on each side whichdevelops just after the curvature; sternal surfaceof abdomen and urites punctate and withpubescence.

Etymology. This new species is named inmemory of the Danish author, critic, architectand designer Poul Henningsen (Ordrup, 9September 1894 - Hillerød, 31 January 1967),in recognition of his cultural contributions.

Syninclusions. A few wood remains and twostellate trichomes.

Remarks. The amber piece measures 9 x 17 x3 mm. The inclusion has damaged antennae, theright one preserved up to the second article (anda small part of the third) and the left onepreserved until the antennomere VI, and also hasthree missing legs from tarsi, or tibiae and tarsi.

Malthodes (s. str.) moellehavei n. sp.(Fig. 16)




Differential diagnosis. Malthodesmoellehavei n. sp. shows intermediate size andcharacters of extant species of both brevicollis-group and minimus-group (Liberti 2016).Concerning the fossil species (Kuśka &Kupryjanowicz 2005; Kuśka & Kania 2010;Fanti 2017ab; Fanti & Vitali 2017), it differsfrom M. ceranowiczae Kuśka &Kupryjanowicz, 2005 in how the last tergite(tg10) is more elongated and with apical tips,and in the more curved last sternite (st9); fromM. michalskii Fanti, 2017 in the longer,


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narrower and more deeply apically incised laststernite and in the longer last tergite withsinuous apex, and in particular, from M. suciniKuśka & Kania, 2010 in the wider penultimatetergite (tg9) and in the apically sinuous lasttergite (tg10). Furthermore, it differs from M.henningseni n. sp. in the tg10 sinuous at apexand st9 apically forked, while it is not forkedbut with lateral appendages near the apex in M.henningseni n. sp.

Description. Adult, winged, slender. Male,based on the last urites modified and the longantennae. Body length 3 mm; antennae 2.1-2.2mm. Entirely dark brown-blackish withoutyellow spots on elytra. Head completelyexposed, rounded, slightly larger than pronotum,equipped with shallow punctation and pubescent.Eyes rounded, convex and prominent. Maxillarypalps 4-segmented, unequal in length, with theterminal one globular - roundish, stout anddistally very pointed. Labial palps 3-segmentedwith the last segment globular and pointed.Antennae filiform, 11-segmented, very long,surpassing the elytra and almost reaching thelast abdominal segments; scape very elongated,not club-shaped and only slightly enlargedapically; pedicel filiform, about one-half as longas the scape; antennomere III as long as thepedicel; antennomeres IV-XI filiform,elongated, subequal in length, slightly longerthan second or third antennomere; allantennomeres with scattered setae. Pronotumsmall, slightly transverse, narrower anteriorly,equipped with dense shallow punctation, basalmargin straight, sides concave and anglesrounded, surface undulate. Scutellum triangular,with very elongated and slightly rounded apex.Elytra slightly wider than pronotum, short,slender, reaching about the abdominal half, wideat base and very narrowed in the apical half, withlargely roundish apex; surface slightly wrinkledand pubescent. Posterior wings wide and long,dark, surpassing the elytra and the abdominalsegments. Legs slender; coxae robust;trochanters elongated; femora enlarged; tibiaecylindrical, very thin, strongly longer thanfemur. Tarsal formula 5-5-5 and tarsomeresrobust; tarsomere I long and more than two

times longer than second; tarsomere III robustand shorter than second; tarsomere IV bilobed;tarsomere V slender; claws simple. Penultimatetergite (tg9) transverse and wide with sidesslightly bent; last tergite (tg10) lobe-shaped andelongated, narrower than tg9, with sinuous apexand apical tips; last sternite (st9) long, stronglycurved, apically deeply forked (V-shaped) withthe lobes narrow and rounded at apex; sternalsurface of abdomen punctate and withpubescence.

Etymology. This new species is named inhonour of the Danish priest, author and lecturerJohannes Volf Møllehave, to thank him for hiswork and to further recognize him for the awardsthat he has earned.

Syninclusions. A few wood remains and onestellate trichome.

Remarks. The rectangular piece of ambermeasures 28 x 11 x 4 mm and the inclusion isperfectly conserved (especially the crucialcharacter of the last tergites and sternites), withthe left antenna aligned along the body and thelegs huddled.

Tribe incertae sedisGenus Kuskaella Fanti & Kupryjanowicz,2017

Kuskaella bajerae n. sp.(Figs. 17 - 18)




Differential diagnosis. This specimen istentatively attributed to the extinct genus


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Kuskaella, based on the habitus, characters andthe very similar long and enlarged apicallyterminal appendix. But in this new species, it isnot completely clear whether this is the laststernite or instead, a part of the aedeagus. Theonly other known species of this genus -Kuskaella macroptera Fanti & Kupryjanowicz,2017 - shows a different length ofantennomeres and particularly, a different shapeof sternites. The penultimate is short andslightly transverse and the last is elongated androundish (Fanti & Kupryjanowicz 2017).

Description. Adult, winged, small and slender.Male, based on the last long sternite and for thevisible part of aedeagus. Body length 3.9 mm;antennal length 3.0 mm. Entirely brown-testaceous. Head exposed, globular, convex, aswide as pronotum, equipped with strongpunctation. Eyes large, roundish, convex.Mandibles not visible. Maxillary palpomeresunequal; first segment very short, secondelongated and globular; third small and short;last globular and equipped with a thin and shortapical appendix. Labial palps 3-segmented withthe last article globular and pointed. Antennae11-segmented, filiform, long, reaching the halfof the elytral length; scape very long, inflated;pedicel robust and more than two times shorterthan scape; antennomere III robust and longerthan pedicel; antennomere IV more robust andlonger than III; antennomeres V-XI, filiform,thinner than others and subequal in length; allantennomeres covered with short and sparsesetae. Pronotum transverse, large at base andnarrowed from half length to apex, with stronglybordered basal angles; apical margin veryslightly curved; basal margin straight; surfaceirregular and equipped with long pubescence andshallow punctation. Scutellum triangular, short,wide and with strongly rounded apex. Elytra long,slender, slightly wider than base of pronotum,and surpassing the last abdominal segments,parallel-sided with rounded apex, equipped withshort setae and rugosity. Posterior wings opaqueand slightly longer than elytra and abdominalsegments. Sternum quadrangular, abdominalsegments strongly transverse and short.Aedeagus (?) with a very elongate lobe, narrow,

and apically enlarged and roundish. Legs robust;coxae massive; trochanteres elongated androundish at apex; femora robust, enlarged; tibiaestout, cylindrical, slightly enlarged apically andslightly longer than femora. Tarsal formula 5-5-5; first tarsomere elongated, widened apicallyand approximately 1.3 times longer than second;second and third tarsomere elongated andwidened apically; fourth short and bilobed; fifthelongated, robust, wide and flat; claws simplewithout denticle.

Etymology. This new species is named inmemory of the Danish women’s rights activistand pacifist Pauline Matilde Theodora Bajer(Frederikseg - Herlufmagle Sogn, 4 January1840 - Copenhagen, 4 March 1934).

Syninclusions. Air bubbles and wood remains.

Remarks. The amber piece is yellow-reddishin colour and measures 27 x 9 x 5 mm. Theinclusion is well visible and almost completeexcept for the lack of an anterior leg and amesothoracic leg conserved until the secondtarsomere. The left antenna is folded and a littlerolled up along the body.

Subfamily Silinae Mulsant, 1862Tribe Silini Mulsant, 1862Genus Autosilis Kazantsev, 2011

Autosilis annisettaekoppelae n. sp.(Fig. 19)




Differential diagnosis. Concerning the fossilrecords of the genus Autosilis, only one


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Fig. 17. Kuskaella bajerae n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 1.0 mm; B:Holotype, ventro-lateral view, bar = 1.0 mm; C: Holotype, detail of last ventrites, bar = 500 μm;D: Holotype, detail of apex of elytra, bar = 200 μm.

Fig. 18. Kuskaella bajerae n. sp. in Baltic amber. Holotype, detail of pronotum, bar = 200 μm.


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fragment strictly related to the extant (cfr.) A.nitidula (Fabricius, 1792) from the Tertiarystrata of Aix-en-Provence, France, is known(Serres 1843; Pictet 1854; Fanti 2017a). Otherremains, such as Silis sp., are known fromEocene Baltic amber (Klebs 1910; Bachofen-Echt 1949; Fanti 2017a), but given the pastconfusion between the two genera, they alsocould be members of Autosilis. One specimenof Silis ruficollis (Fabricius, 1775) is alsoknown from the Holocene (5200-5100 y/o)deposit of Sweet Track, United Kingdom(Girling 1979). The new species here describedshows a coloration (dark brown) different fromthat of A. nitidula, which is completely blackin males and black with orange-red pronotumin females. The genus Autosilis is characterizedby glabrous pronotum in the anterior half andwithout deep cavities (Kazantsev 1994).

Description. Adult, winged. Male, defined onthe basis of pronotum with modified sides.Body length ~8 mm but difficult to be certain

because the head is folded and unobtrusive.Entirely dark brown. Head small and roundish.Eyes subspherical. Maxillary palpomeresunequal; first segment very short; secondelongated; third with intermediate lengthbetween the first and the second; last segmentsecuriform. Labial palps 3-segmented. Antennaeinserted far away from the eyes, slightlysurpassing the metafemurs, 11-segmented,filiform, covered with small setae; scapeglobular, massive and elongated; pedicelextremely short; antennomere III twice longerthan pedicel; antennomeres IV-VI subequalelongated with small expansion at sides;antennomeres VII-XI filiform, very elongated.Pronotum rounded, smooth, without deepcavities, flat except for the basal part which isslightly concave, with the lower process at sidesvery large and the upper process at sides verythin, slightly pointed and about one-half as longas the lower process. Scutellum very wide atbase and elongated at apex, which is roundish.Elytra as wide as pronotum (considering the end

Fig. 19. Autosilis annisettaekoppelae n. sp. in Baltic amber. A: Holotype, dorsal view, bar = 1.0mm; B: Holotype, ventral view, bar = 1.0 mm; C: Holotype, lateral view, bar = 1.0 mm; D: Holotype,detail of scutellum and humeral zone of elytra, bar = 500 μm.


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of the lower process) with microsculpturerugose and some setae, elongated and coveringthe last abdominal segments; apex stronglyrounded. Posterior wings covered by elytra.Sternum and abdominal segments covered withnumerous setae. Legs short and pubescent;coxae roundish; trochanters elongated andtriangular with roundish apex; femora veryrobust and massive; tibiae short as femora,cylindrical, thin and with a spur at apex. Tarsalformula 5-5-5; first segment elongated andslightly enlarged apically; second shorter thanfirst; third segment subequal to second; fourthsegment very bilobed at sides; fifth veryelongated; claws simple with tooth at base onthe inner claws.

Etymology. This new species is named inhonour of the Danish singer Annisette Koppel(born Hansen), in recognition of her longcareer.

Syninclusions. Air bubbles, several woodremains (particles), pyrite.

Remarks. The amber piece measures 10 x 28 x5 mm and shows numerous internal cracks. Theinclusion is probably complete but ventrally, itis covered by a white cotton-like cloud(emulsion, milky substance), which does notallow a good view of some parts. In fact, thehead is folded and the left antenna is partiallyvisible.

DISCUSSION

Eridanula n. gen. and Noergaardia n. gen., aswell as Sucinocantharis Kuśka & Kania, 2010and Cacomorphocerus Schaufuss, 1892, areclearly strictly related to the same lineage andtherefore, all belong to the recently establishedtr ibe Cacomorphocerini (Fanti &Kupryjanowicz 2018). This currentlycompletely extinct tribe is characterised bysaucer-shaped “central” and supernumerary (12to 19) antennomeres (Fanti & Kupryjanowicz2018). In contrast, Palmnickeneoceras n. gen.is assigned to the tribe Cantharini for showing

11 antennomeres, little dilated and very slightlysaucer-shaped, while Juratelacrima n. gen. onthe other hand, it has also an aspect similar tosome Podabrini Gistel.

Cantharis seems rare in amber, with only threespecies (included C. mikkelsenorum n. sp.)known until now (Fanti 2017a). This is probablyrelated to their quite large body size but also tothe fact that this genus usually visits and eatsupon flowers, while Malthodes and Malthinussearch for their prey on leaves of trees andbushes and thus are more easily trapped in theresin. The body size of these last genera isminute, generally below 3-4 mm (to ~5.5 mmin Malthinus), so that 12 species of Malthodes(included M. henningseni n. sp. and M.moellehavei n . sp.) and two Malthinus(included M. rifbjergi n. sp.) are known in Balticamber, Rovno amber and Brunstatt brown coalsof Alsace (Förster 1891; Kuśka &Kupryjanowicz 2005; Kazantsev 2010; Kuśka& Kania 2010; Kazantsev & Perkovsky 2014;Fanti 2017b; Fanti & Vitali 2017; Fanti &Michalski 2018). The body size could alsoinfluence the difficulty to find members of thegenus Rhagonycha Eschscholtz.

Podistra (Absidia) kloevedali n. sp. is the firstspecies of this genus described as a fossil; infact, previously, Podistra was known only atgeneric level (Klebs 1910; Bachofen-Echt1949; Fanti 2017a). The genus usually lives onmountains, and currently, only three extantspecies of the subgenus Absidia are known(Kazantsev & Brancucci 2007).

The discovery of two species of the genusThemus (Th. pristinus and Th. bennyiandersenin. sp.) in Baltic amber allows some assumptionsto be made about the origin of the genus and theage of the amber. Currently, Themus iswidespread in Iran, Kyrgyzstan, Tajikistan,Pakistan, Afghanistan, India, Nepal, Bhutan,China, Kuril, Japan, Indochina, Sumatra, Java andBorneo (Švihla 2008; Fanti & Ghahari 2016)and is not present in Europe (Kazantsev &Brancucci 2007). Because Europe was


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separated from Asia by the Turgai Strait-TurgaiSea during the Eocene, it is possible thatThemus is older and originated in theCretaceous or the Early Paleocene, whenEurope and Asia where connected. Otherwise,if the Baltic amber dates back to Eocene,Themus reached Asia in the Oligocene followingthe climate cooling due to the draining of theTurgay Sea. The other possible hypothesis is thatthe Baltic amber is more recent, dating back tothe Oligocene, and that Themus reached Europefrom Asia.

The attribution of Kuskaella bajerae n. sp. tothe subfamily Malthininae is clear from thehabitus, the globular and pointed last maxillarypalpomere and various other characters.Nonetheless, the mandibles are not visible andthe tribe remains incertae sedis (Fanti &Kupryjanowicz 2017).

The subfamily Silinae is rare in Europe, whereonly the genera Silis and Autosilis are present,while it is particularly numerous in Asia andAmerica, and seems to be very diversified inthe Baltic Eocene.

The Anders Damgaard amber collection, alsocontains other soldier beetles, such as twospecimens of Cacomorphocerus sp.(ALDC0056/ALD.Ba.Can.7 and ALDC0058/ALD.Ba.Can.9), whose photos were shown inFanti (2017a), and a female of Malthodes sp.(ALDC0051/ALD.Ba.Can.2).

ACKNOWLEDGEMENTS

We are indebted to Francesco Vitali(Luxembourg), for the precious and veryconstructive advices that have enhanced themanuscript, and to Alessio Morelli (Pianella,Pescara, Italy) for the excellent drawings. Weare particularly grateful to Mary Pankowski(Rockville, Maryland, USA) for theimprovement and editing the article.

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Received: 27.04.2018.Accepted:30.07.2018.

Published: 10.08.2018.


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